A test of the "dear enemy effect" in the strawberry dart-poison frog (<Emphasis Type="Italic">Dendrobates pumilio</Emphasis>)

Some territorial animals exhibit a form of social recognition, commonly termed the "dear enemy effect", in which territory residents display lower levels of aggression toward familiar neighbors compared to unfamiliar individuals who are non-territorial "floaters". Despite the widespread occurrence of territorial social systems and use of acoustic signals for communication in anuran amphibians, only two previous studies have demonstrated vocally mediated dear enemy behavior in a territorial frog. In this study, I conducted neighbor-stranger discrimination playback experiments in a third species of territorial frog, the strawberry dart-poison frog, Dendrobates pumilio (Anura, Dendrobatidae). In the first experiment (n=24), I broadcast the calls of a subject's nearest neighbor and the calls of an unfamiliar individual from the approximate midpoint between the subject's and the neighbor's territories. Although males responded to the stimuli, they did not exhibit differential responses to the calls of neighbors and strangers. In a second experiment (n=22), I broadcast the calls of a neighbor and a stranger to subjects through a speaker located in the approximate center of the neighbor's territory. Males also responded to the playback, although less intensely than in the first experiment, but no discrimination between the calls of neighbors and strangers was found. Thus, territorial males of the strawberry dart-poison frog appear not to discriminate behaviorally between the advertisement calls of neighbors and strangers. Several proximate and ultimate-level hypotheses for this lack of vocally mediated neighbor-stranger discrimination are discussed.Communicated by T. Czeschlik     

Heart rate changes reveal that little blue penguin chicks (Eudyptula minor) can use vocal signatures to discriminate familiar from unfamiliar chicks

Researchers investigating social recognition typically measure only behavioural responses during discrimination tests - physiological changes have been largely ignored. We examined whether little blue penguin chicks (Eudyptula minor) could distinguish siblings from other chicks using auditory cues, by measuring behavioural and heart rate changes during playback experiments. Chicks were exposed to five treatments: the begging calls of siblings, neighbouring chicks and unfamiliar chicks, and two controls (heterospecific begging calls and music). We also determined if chicks developed distinctive begging calls, by using F-ratios to quantify inter- versus intra-individual variability in a range of acoustic parameters, and applying a discriminant-function analysis. Inter-individual variation was greater for pitch than for temporal or amplitude parameters, suggesting that call pitch may be important for individual recognition. The discriminant-function analysis showed each chick's calls were distinctive and could act as a vocal signature. Treatments did not instigate different behavioural responses. However, chick heart rates during playback of sibling calls were significantly higher than those recorded during stranger, but not neighbour, playback. A simple recognition system based on familiarity may allow this plesiomorphic and loosely colonial penguin to gain at least some of the benefits associated with more advanced sibling recognition systems (some highly colonial seabirds discriminate siblings from neighbouring chicks). Heart rate could be a useful measure of social recognition abilities, particularly in species where changes in behaviour are not always evident or are difficult to observe.     

Loud calls as a mechanism of social coordination in a fission–fusion taxon, the white-bellied spider monkey (Ateles belzebuth)

Spider monkeys (Ateles spp.) live in social groups that exhibit high levels of fission–fusion dynamics, in which group members form subgroups of varying sizes and compositions. Within these fluid societies, how individuals establish contact with dispersed group members with whom they might choose to associate remains unclear. Long-range vocalizations might facilitate interactions between group members and provide a means of social coordination in fission–fusion societies. We evaluated this possibility for one spider monkey vocalization, the loud call, by examining calling behavior, the relationship between loud calls and changes in subgroup size, and the response of individuals to distant calls and playback experiments in a single study group. We found that 82 % of loud calls were emitted within 30 min of a call from a different location, suggesting that individuals frequently emit loud calls in response to the calls of distant group members. Subgroups that emitted loud calls, especially those that responded to distant calls, were much more likely to experience an increase in subgroup size within an hour after calling than those that did not. Animals also approached distant loud calls more than they avoided or ignored these calls. Finally, playbacks of male calls demonstrated that females respond preferentially to the calls of some individuals over others. Taken together, these results provide support for the hypothesis that spider monkey loud calls function to facilitate and initiate interactions between dispersed group members and suggest that vocal signals can play an important role in influencing social interactions in fission–fusion societies.     

Playback and speaker-replacement experiments on song-based neighbor,stranger, and self discrimination in male Red-winged Blackbirds

Summary Two experiments were conducted to deterine whether territorial male Red-winged Blackbirds discriminate among neighbor, stranger, and self songs. In the first experiment, recordings of the three classes of songs were played to territorial males. Responses to playback of stranger song, as measured by song rate, intensity of Song Spread display, and closest approach, were significantly stronger than to playback of neighbor song. In addition, stranger song elicited significantly more intense Song Spread displays than did self song.In the second experiment, territorial males were removed from their territories and each was replaced with speakers broadcasting recorded songs of a stranger, and of the removed male. Broadcasts of the removed male's own songs were more effective in discouraging trespassing by neighboring male redwings than were broadcasts of songs of a stranger. However, the two broadcasts did not differ in their ability to deter nonneighboring male redwings from flying through or trespassing within the speaker territory.We suggest that neighbor-stranger discrimination in the Red-winged Blackbird develops as a consequence of associative learing.     

Early learning of discrete call variants in red crossbills: implications for reliable signaling

The identification of appropriate companions and mates is essential to both speciation and the maintenance of species through prezygotic isolation. In many birds, social assortment is mediated by vocalizations learned through imitation. When imitative vocal learning occurs throughout life, emergent shared signals reflect current social associations. However, when vocal and genetic variation arises among populations, shared learned signal variants have a potential to reflect cultural or genetic origin and to limit social and reproductive intermixing, provided that signal learning occurs prior to dispersal. The red crossbill (Loxia curvirostra) is a bird species in which discrete contact call variants are associated with morphological variation, raising the possibility that learned calls play a role in limiting intermixing. I examined the process of early call learning to determine if contact call variants have a potential to limit intermixing in crossbills. I conducted a captive playback study to nestlings to evaluate potential learning predispositions. I also cross-fostered nestlings to adoptive adult pairs of either their own or a different call variant than their biological parents to assess the degree of vocal learning plasticity. Results show that young crossbills imitate the call structures of adoptive parents, generating shared family-specific calls, which could facilitate family cohesion. Learning processes that generate family-specific calls could also ensure that discrete call variants are transmitted across generations, making call variants reliable signals of crossbills’ morphological and genetic backgrounds.     

Chick-a-dee call syntax,social context,and season affect vocal responses of Carolina chickadees (<Emphasis Type="Italic">Poecile carolinensis</Emphasis>)

Chick-a-dee calls in many chickadee (Poecile) species are common vocal signals used in a diversity of social contacts. The calls consist of four notes, A, B, C, and D, which follow simple rules of syntax (note ordering and composition) to generate many unique call types. We used field playbacks with Carolina chickadees, P. carolinensis, to ask whether violations of a syntactical rule affected their vocal responses. We show that chickadee responses to typical calls (e.g. AAAACCCC and CCCCDDDD) differ from responses to atypical calls (e.g. CACACACA and DCDCDCDC) depending on playback note composition, season, and social context (presence of heterospecifics). In the fall/winter, playbacks of typical calls with A and C notes elicited the greatest number of A and B notes in chick-a-dee call responses and typical calls with D notes elicited the greatest number of C notes, when in the presence of heterospecifics. In contrast, the corresponding atypical calls did not elicit similar responses. This suggests communicative significance is lost in calls that violate a rule of syntax in the fall/winter. In the spring, neither chickadee feebeefeebay song rate nor chick-a-dee calls responses differed by playback type. We suggest that call syntax is less salient for mated pairs in the spring than it is for fall/winter flocks that rely more on conspecific communication for foraging success and flock cohesion. This study represents the first experimental evidence that chickadees attend to both note composition and ordering in chick-a-dee calls.Communicated by W.A. Searcy     

Anti-predator behavior of group-living Malagasy primates: mixed evidence for a referential alarm call system

Many mammals warn conspecifics with alarm calls about detected predators. These alarm calls are either functionally referential, urgency based, or they can have multiple functions, including predator deterrence. The taxonomic distribution of these alarm call systems is uneven, with primates providing the best-known examples for a functionally referential system and rodents most examples of an urgency-based system. Reports of different alarm call systems in lemurid primates prompted us to examine the anti-predator behavior of two additional lemur species. In an experimental field study we exposed adult redfronted lemurs (Eulemur fulvus rufus) and white sifakas (Propithecus verreauxi verreauxi) to playbacks of vocalizations of their main aerial and terrestrial predators, as well as to their own alarm calls given in response to the presentation of these predators. We scored the subjects' immediate behavioral responses, including alarm calls, from video recordings made during the first minute following a playback. We found that both species gave specific alarm calls only in response to raptor playbacks and the corresponding alarm calls, whereas calls given in response to carnivores and the corresponding alarm calls were also observed in other situations characterized by high arousal. Other behavioral responses, such as gaze and escape directions, corresponded to the hunting strategies of the two predator classes, suggesting that the corresponding vocalizations were categorized correctly. These two lemur species, which represent different families, have therefore independently evolved a mixed alarm call system, characterized by functionally referential calls for diurnal raptors, but not for carnivores. Electronic supplementary material to this paper can be obtained by using the Springer LINK server located at http://dx.doi.org/10.1007/s00265-001-0436-0 Electronic Publication     

Effects of multiple-speaker playbacks on aggressive calling behavior in the treefrog <Emphasis Type="Italic">Dendropsophus ebraccatus</Emphasis>

In addition to producing signals, males of chorusing species also act as receivers by adjusting properties of their vocalizations in response to those of other nearby individuals. Although it is likely that males are responsive to more than one other individual, most playback studies investigating male response have involved dyads in which vocal responses are measured to stimuli presented from a single speaker. In this study, I explored changes in both the propensity to give aggressive calls and the temporal properties of those calls in response to the playback of multiple aggressive call stimuli in the treefrog Dendropsophus ebraccatus. I found that males were sensitive to both the number of simulated aggressive callers and their specific call characteristics. Males generally gave a highly aggressive response to the first stimulus presented, but their response to the modification of this stimulus by the addition or subtraction of a simulated competitor depended on the degree of aggressiveness of the stimuli. Males tended to decrease their aggressive responses when either a more aggressive call was silenced or a less aggressive call was added and to increase their aggressive responses in the opposite situation. Aggressive calling in this species is clearly affected by complex changes in the social environment and I suggest that future studies explore these issues in other species to improve the understanding of communication interactions.     

Neighbor–stranger discrimination in Audubon's shearwater (Puffinus l. lherminieri) explained by a “real enemy” effect

Neighbor–stranger discrimination (NSD) occurs when animals respond with more aggression to strangers than to territorial neighbors. NSD has been reported in many species that defend multi-purpose territories for breeding and foraging, but it is rare among species that defend other types of territories. For birds that defend only their nest sites, there is no experimental evidence for NSD, and observational studies have provided mixed results. In a colony of Audubon's shearwaters (Puffinus l. lherminieri), I played back the calls of a neighbor and a stranger to males defending nest sites. Subjects responded with longer calls to playbacks of strangers than to those of neighbors. In shearwater colonies, strangers are often birds looking for future breeding sites. In contrast, there is no evidence that established breeders compete with their neighbors for any resources. Shearwaters should benefit from NSD because strangers represent a “real enemy” and established neighbors do not.     

Sound localization by monkeys: A field experiment

Summary Characteristic responses shown by free ranging mangabeys to playback of species-specific vocalizations allow field measurement of their sound-localizing abilities. The median error of localization of single 10 to 15 s calls heard through several hundred meters of tropical forest was only 6°.     

Discrimination of different social companions in spectacled parrotlets (Forpus conspicillatus): evidence for individual vocal recognition

Individual recognition is generally assumed to be a prerequisite for establishing and maintaining a complex social system. Indeed, there is good evidence that highly social species have complex systems of vocal communication with individual recognition by acoustic cues. In this study, we provide experimental evidence that vocal class and individual recognition is present in a non-passerine bird, the spectacled parrotlet (Forpus conspicillatus). Spectacled parrotlets live in a complex system of social relationships. Soon after fledging, the young establish close sibling relationships which are important for successful socialization, pairing and reproduction. In a series of playback experiments we tested if spectacled parrotlets use contact calls for vocal recognition. The results showed that spectacled parrotlets discriminate between the contact calls of different social categories. Adult birds preferred to respond to the contact calls of their mates. Subadult individuals recognized the contact calls of their siblings. During the period of pair bond formation, the affiliative contacts to the siblings decrease, but the parrotlets continue to respond to the calls of their siblings. This is the first evidence that vocal sibling recognition might outlast the period of strong sibling interaction and extends into the period of pair bond formation. In cases of mate loss or divorce, the acoustic contact to their siblings might facilitate the re-establishment of close sibling relationships. Received: 29 October 1997 / Accepted after revision: 5 April 1998     

Are vocal signals used to recognize individuals during male–male competition in greater prairie-chickens (Tympanuchus cupido)?

Vocal signaling can be an important component of vertebrate communication during social interactions. If vocalizations vary among individuals but are consistent within a given individual, they may be used to discriminate among individuals. In many species, territorial males use vocalizations to discriminate between neighbors and strangers and either respond more aggressively toward strangers relative to neighbors (“dear enemy” effect) or they respond more aggressively toward neighbors relative to strangers (“nasty neighbor” effect). In the greater prairie-chicken (Tympanuchus cupido), male vocalizations are an integral part of the display males produce on leks. We investigated whether male greater prairie-chickens discriminate among familiar individuals on their own territory, familiar individuals outside their normal territory and strangers from a nearby lek. Vocal characteristics varied among males, suggesting that vocalizations may potentially be used by prairie-chickens to identify individuals. Males responded to playback of prairie-chicken calls by vocalizing at a faster rate and approaching the playback speaker, but did not vary in their response to the vocalizations based on the identity of the caller. Our results suggest that variation is present among the vocalizations of individual male greater prairie-chickens, but males do not appear to discriminate among familiar individuals and strangers based solely on their “boom” vocalizations. Greater prairie-chicken vocalization likely functions as a way of announcing that a territory is occupied and defended, but it may also serve as a way of advertising to conspecifics or as a signal that is secondary to other forms of communication.     

Greater spear-nosed bats give group-distinctive calls

Individually distinctive vocalizations are ubiquitous; however, group distinctive calls have rarely been demonstrated. Under some conditions, selection should favor calls indicating social group membership in animals that forage in groups. Greater spear-nosed bats (Phyllostomus hastatus) give calls that appear to facilitate recognition of social group mates who are unrelated. Females give loud broadband (4–18 kHz) vocalizations termed screech calls when departing on foraging trips and at foraging sites. Screech calls help to establish foraging groups among social group members, and to maintain contact over the long distances they travel while foraging. I test two hypotheses about how screech calls may be structured to convey caller identity. Individual calls may be distinct and group members may learn to recognize each individual's calls and to associate the individual with the social group. Alternatively, groups may give distinct calls and individuals within groups may share call characteristics. To test these hypotheses I conducted multivariate acoustic analysis of multiple calls from 28 bats from three social groups. Although the ubiquity of individually distinctive calls in other taxa makes this result more likely, the results reveal that group calls are highly distinctive. Individual bats within groups are statistically indistinguishable. Calls appear to decrease slightly in frequency as bats age. Call convergence among unrelated group mates implies vocal learning in this species. Received: 28 March 1996 / Accepted after revision: 6 October 1996     

Similar is not the same: Social calls of conspecifics are more effective in attracting wild bats to day roosts than those of other bat species

Many bat species regularly need to find new day roosts as they require numerous shelters each breeding season. It has been shown that bats exchange information about roosts among colony members, and use echolocation and social calls of conspecifics in order to find roosts. However, it is unclear if wild bats discriminate between social calls of conspecifics and other bat species while searching for roosts. Furthermore, the extent that bats are attracted to potential roosts by each of these two call types is unknown. We present a field experiment showing that social calls of conspecifics and other bat species both attract bats to roosts. During two summers, we played back social calls of Bechstein’s bats (Myotis bechsteinii) and Natterer’s bats (Myotis nattereri) from different bat boxes that can serve as roosts for these species. All experimental bat boxes were monitored with infrared video to identify the approaching bat species. Three species (M. bechsteinii, M. nattereri, and Plecotus auritus) approached the boxes significantly more often during nights when bat calls were played compared to nights without playbacks. Bechstein’s bats and Natterer’s bats were both more attracted to social calls of conspecifics than of the other species, whereas P. auritus did not discriminate between calls of either Myotis species. Only Bechstein’s bats entered experimental boxes and only at times when calls from conspecifics were played. Our findings show that wild bats discriminate between social calls of conspecifics and other bat species although they respond to both call types when searching for new roosts.     

Individual specific contact calls of pallid bats (<Emphasis Type="Italic">Antrozous pallidus</Emphasis>) attract conspecifics at roosting sites

Contact calls are utilized by several bird and mammal species to maintain group cohesion and coordinate group movement. From a signal design perspective, contact calls typically exhibit acoustic features that make them easily localizable and encode information about individual or group identity. Pallid bats (Antrozous pallidus) are unusual among vespertilionids in that they often emit a loud, partially audible frequency-modulated social call several times in rapid succession while in flight. This call appears to function as a contact call in that it is frequently given when bats return from foraging and perform circular flights before entering a crevice roost. However, the degree to which pallid bats respond to the calls of conspecifics and what information is provided in the call is unknown. Thus, the goal of this study was to investigate pallid bat calling behavior to determine if calls attract roostmates or elicit responses from them and provide sufficient information for individual recognition. In playback studies, we found that contact calls, elicit calls, and approaches and that free-flying bats respond more to familiar than unfamiliar calls. In addition, analysis of frequency and temporal measurements of calls collected from multiple sites and spectral cross correlation analysis of calls recorded from the same radio-tagged bats on multiple evenings revealed that the frequency pattern of contact calls is highly repeatable over time within individuals but exhibits significant differences among individuals. Thus, contact call structure appears to be unique to individuals and stable through time, which makes these calls well-suited for roostmate recognition.     

A test of multiple hypotheses for the function of call sharing in female budgerigars, Melopsittacus undulatus

In many social species group, members share acoustically similar calls. Functional hypotheses have been proposed for call sharing, but previous studies have been limited by an inability to distinguish among these hypotheses. We examined the function of vocal sharing in female budgerigars with a two-part experimental design that allowed us to distinguish between two functional hypotheses. The social association hypothesis proposes that shared calls help animals mediate affiliative and aggressive interactions, while the password hypothesis proposes that shared calls allow animals to distinguish group identity and exclude nonmembers. We also tested the labeling hypothesis, a mechanistic explanation which proposes that shared calls are used to address specific individuals within the sender–receiver relationship. We tested the social association hypothesis by creating four–member flocks of unfamiliar female budgerigars (Melopsittacus undulatus) and then monitoring the birds’ calls, social behaviors, and stress levels via fecal glucocorticoid metabolites. We tested the password hypothesis by moving immigrants into established social groups. To test the labeling hypothesis, we conducted additional recording sessions in which individuals were paired with different group members. The social association hypothesis was supported by the development of multiple shared call types in each cage and a correlation between the number of shared call types and the number of aggressive interactions between pairs of birds. We also found support for calls serving as a labeling mechanism using discriminant function analysis with a permutation procedure. Our results did not support the password hypothesis, as there was no difference in stress or directed behaviors between immigrant and control birds.     

Individual recognition of neighbors by song in a suboscine bird, the alder flycatcher Empidonax alnorum

The ability of territorial males to discriminate between songs of their neighbors and songs of strangers has been demonstrated in 27 species of songbirds. Such experiments test only the ability of a subject to discriminate between two classes of stimuli, familiar (neighbors) and unfamiliar (strangers) songs. Individual recognition of neighbors is a finer, more complex type of discrimination. The ability of territorial males to recognize individual neighbors by song has been documented in 12 species of oscine passerines (Passeriformes, Passeri), but has never been demonstrated in suboscine passerines (Tyranni). We investigated recognition of songs of individual neighbors in a suboscine, the alder flycatcher (Empidonax alnorum). We performed a series of song playback experiments and recorded responses of territorial males to songs of neighbors and songs of strangers broadcast from two locations, the neighbor boundary and an opposite boundary. Subjects responded more aggressively to songs of a neighbor when played from the opposite boundary than when played from the neighbor boundary. They responded with equal aggression to songs of strangers regardless of location of playback. The difference in response to neighbor songs between speaker locations and the lack of a difference in response to stranger songs indicate that territorial males associate a particular song with a particular location (territory), and thus recognize individual neighbors.Communicated by I. Hartley     

Facilitating climate‐change‐induced range shifts across continental land‐use barriers

Climate changes impose requirements for many species to shift their ranges to remain within environmentally tolerable areas, but near‐continuous regions of intense human land use stretching across continental extents diminish dispersal prospects for many species. We reviewed the impact of habitat loss and fragmentation on species’ abilities to track changing climates and existing plans to facilitate species dispersal in response to climate change through regions of intensive land uses, drawing on examples from North America and elsewhere. We identified an emerging analytical framework that accounts for variation in species' dispersal capacities relative to both the pace of climate change and habitat availability. Habitat loss and fragmentation hinder climate change tracking, particularly for specialists, by impeding both propagule dispersal and population growth. This framework can be used to identify prospective modern‐era climatic refugia, where the pace of climate change has been slower than surrounding areas, that are defined relative to individual species' needs. The framework also underscores the importance of identifying and managing dispersal pathways or corridors through semi‐continental land use barriers that can benefit many species simultaneously. These emerging strategies to facilitate range shifts must account for uncertainties around population adaptation to local environmental conditions. Accounting for uncertainties in climate change and dispersal capabilities among species and expanding biological monitoring programs within an adaptive management paradigm are vital strategies that will improve species' capacities to track rapidly shifting climatic conditions across landscapes dominated by intensive human land use.     

Paternity alone does not predict long-term investment in juveniles by male baboons

Adult male chacma baboons (Papio hamadryas ursinus) form preferential associations, or friendships, with particular lactating females. Males exhibit high levels of affiliative contact with their friends’ infants and defend them from potentially infanticidal attacks (Palombit et al. 1997). Little is known about males’ associations with juveniles once they have passed the period of infanticidal risk. We conducted an observational, experimental, and genetic study of adult male and juvenile chacma baboons in the Moremi Reserve, Botswana. We identified preferential associations between males and juveniles and used behavioral data and a playback experiment to explore whether those associations have potential fitness benefits for juveniles. We determined whether males preferentially invest in care of their own offspring. We also determined how often males invest in care of their former friends’ offspring. The majority of juveniles exhibited preferential associations with one or two males, who had almost always been their mother’s friend during infancy. However, in only a subset of these relationships was the male the actual father, in part because many fathers died or disappeared before their offspring were weaned. Male caretakers intervened on behalf of their juvenile associates in social conflicts more often than they intervened on behalf of unconnected juveniles, and they did not appear to differentiate between genetic offspring and unrelated associates. Playbacks of juveniles’ distress calls elicited a stronger response from their caretakers than from control males. Chacma males may provide care to unrelated offspring of former friends because the costs associated with such care are low compared with the potentially high fitness costs of refusing aid to a juvenile who is a possible offspring.     

Social dominance modifies behavioral rhythm in a queenless ant

Social insects provide an intriguing model system in chronobiology. Typically, an egg-laying queen exhibits arrhythmicity in activity while foraging worker has clear rhythmicity. In the queenless ant, Diacamma sp., from Japan, colony members lack morphological caste, and reproductive differentiation occurs as a consequence of dominance hierarchy formation. Their specialized dominance interaction “gemmae mutilation”, provide us a fascinating model system to investigate the effect of social dominance on rhythmic ontogeny. Measurement of individual rhythms revealed that they have clear circadian rhythm at eclosion but it is diminished by social mutilation of gemmae. Moreover, unlike highly eusocial species, mated egg-layer (i.e., gamergate) possessed a circadian rhythm even after mating in Diacamma. Measurement of colony-level rhythms revealed that gemmae mutilations are performed in the limited time of the day, but foraging occurs around-the-clock. The above finding is a novel form of temporal organization in social insects, providing a new insight in morphologically casteless species. We discuss the causes and consequences of rhythmic variability in social organization.